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IIBRAHY 

OF THE. 

UN IVER.SITY 

Of ILLINOIS 

590.5 
FI 

v. 37-38 



BIOLOtii 



Latest Date stamped below. 




L161 O-1096 



A Revision of the Tribe Amblyopinini: 

Staphylinid Beetles Parasitic on Mammals 

Charles H. See vers 
Research Associate, Division of Insects 

INTRODUCTION 

This paper 1 is concerned with a group of staphylinid beetles 
that live as ectoparasites on rodents and marsupials. These staphy- 
linids are of unusual interest inasmuch as obligate ectoparasitism is 
a very rare trait among beetles. 

In an earlier paper (Seevers, 1944) I proposed that these ecto- 
parasitic genera be placed in a new subfamily, Amblyopininae, as 
their former position in the Habrocerinae was not a natural one. 
Since that paper was published I have been able to study much 
more material, including specimens of the Tasmanian genus Myo- 
typhlus. It is now evident that the group is morphologically linked 
to the tribe Quediini of the Staphylininae. The genus Myotyphlus, 
in particular, is very close to the Quediini and is doubtfully separable 
on structural grounds. In this revision the group is placed in the 
subfamily Staphylininae as the tribe Amblyopinini. The Neotropical 
species of Amblyopinini are morphologically specialized to the degree 
that separation from the Quediini is not difficult to justify. The 
tribe as a whole may well be justified on the basis of the distinctive 
ecology and behavior of its members. The evolution of obligate 
ectoparasitism in the Staphylinidae must have required genetic 
changes comparable to, if not greater than, those necessary to bring 
about morphological characters generally recognized as separating 
higher categories of Staphylinidae. To group these genera with the 
Quediini obscures their distinctive ecological and physiological adap- 
tations; yet to separate them need not obscure their probable origin 
from quediine stock. 

1 This project was completed during the tenure of a grant in aid from the 
National Science Foundation. 

211 



212 FIELD IANA: ZOOLOGY, VOLUME 37 

A revision of the tribe has been made necessary by the large 
number of new species that have been collected in recent years. 
The size of the group is more than doubled by the addition of 
twenty-three new species. One of the surprising revelations of my 
recent studies is the high degree of host specificity shown by these 
species. While it is true that many of the host records reflect only 
a single capture, there are enough records based on more than one 
to indicate that these beetles have narrowly limited host preferences. 
The idea that the tribe Amblyopinini contains a few, widely dis- 
tributed species must be dismissed. 

It is desirable at this time to revise our concepts of some Neo- 
tropical genera. The limits of Amblyopinus are herein restricted 
somewhat by the separation of two distinctive species groups as new 
genera, Megamblyopinus and Amblyopinodes. 

The distribution records of the genera and species of Amblyo- 
pinini clearly reveal that the group occurs almost exclusively in the 
subtropical and temperate zones of the Neotropical region (see 
distribution map, fig. 28). Mr. Philip Hershkovitz was impressed 
by the fact that these beetles were not present on mammals collected 
by him in the lowlands of Colombia (see notes from his records 
elsewhere in the paper). The records indicate the occurrence of 
Amblyopinini in the Andes of Colombia, Ecuador, and Peru (doubt- 
less in Venezuela, Bolivia and Chile as well) ; the mountains of Guate- 
mala (and almost certainly in other Central American countries and 
Mexico); Mount Roraima in British Guiana; the subtropical zones 
of Rio de Janeiro, Sao Paulo, and Santa Catharina of Brazil; northern 
Argentina and Chile; and the south temperate region of Chile. 

ECOLOGICAL CONSIDERATIONS 

There is no doubt that the Amblyopinini are obligatory ecto- 
parasites. From the time of the first report (Solsky, 1875) the records 
are consistent with regard to this point. Jelski, the first collector, 
observed that Amblyopinus jelskii was firmly attached to the skin 
of living mice just in front of the bases of their tails; as a result 
the hair was eroded in this area and the swollen skin exuded a serous 
secretion. Fauvel (1900) reported the observations of Philippi 
relating to the habits of Edrabius philippianus in Chile. The adults 
and larvae lived exclusively around the anus of Ctenomys and caused 
the same condition of the skin that Jelski had noted. More re- 
cently, Zikan (1939) concluded that Amblyopinus henseli Kolbe is a 
blood-sucking parasite. He noted that the beetles imbedded their 




Fig. 28. Map showing the distribution of the Amblyopinini in South America. 



213 



214 FIELDIANA: ZOOLOGY, VOLUME 37 

mandibles so deeply into the skin that they were difficult to remove 
and that they probably fed on body fluids. Zikan presented no 
direct evidence that Amblyopinus is a blood-sucking parasite, so 
the question is still open. 

With regard to the habits of the species reported in this paper, 
there are interesting field notes made by Philip Hershkovitz while 
on the Chicago Natural History Museum Colombian Zoological 
Expedition (1948-52). The following account is from Mr. Hersh- 
kovitz' field notes: "Ectoparasitic staphylinids were first encountered 
by me in the Valdivia region of Antioquia in the subtropical zone 
of the central Andes of Colombia at altitudes between 4,550 and 
6,500 feet. Beetles found on or near the dead bodies of trapped 
mice gave the impression of being merely free-living scavengers 
or carrion feeders. However, two live individuals of the white- 
throated mouse, Oryzomys albigularis Tomes, with associated 
beetles were kept in a small wire cage and two beetles on one and 
three on the other were observed at intervals during the course of a 
few days. When feeding or at rest, the beetles 'attached' them- 
selves, seemingly by hooking their mandibles onto the fur or skin in 
the angle between the ear and the head. Their bodies were main- 
tained fairly motionless or with intermittent, slow undulatory 
motions, with the long axis at right angles to the point of 'attach- 
ment.' At times all the beetles would congregate behind one of the 
mouse's ears and at other times they could be found behind both 
ears. No interchange of beetles from one mouse to another was 
observed, although this probably took place. When the mice were 
especially active or were disturbed by prodding or handling, the 
beetles 'detached' themselves and scurried or wriggled with remarkable 
agility through the fur. Surprisingly, the hosts appeared to be 
insensible to the movements of the beetles over their bodies and 
across such sensitive parts of the face as the eyeballs and bases of 
the vibrissae. When the hosts relaxed, the beetles returned to their 
position of attachment behind the ears. 

"I was not able to determine the substance of the host that was 
consumed by the parasites. No marks attributable to the beetles 
could be detected. My impression is that the staphylinids may 
seek certain epidermal secretions or incrustations. Like fleas, they 
leave the dead host when the body begins to cool but do not wander 
far. I have found the beetles on the under side of traps that held 
dead mice and under the surrounding vegetation. The beetles are 
comparatively sluggish when detached from their hosts and are easily 
captured by hand. As they seek to avoid light their greatest move- 



SEEVERS: REVISION OF AMBLYOPININI 215 

ments are made when they are exposed to view. On the warm host 
they are remarkably swift and adroit in slipping between one's 
fingers. 

"The beetles were observed on specimens of several small rodent 
species captured in the subtropical zone of the Valdivia region. 
The same species occur in the tropical lowlands of Colombia but 
I have never found these parasites associated with them. Indeed, 
the staphylinids were not found with any mammals captured by 
me in the lowlands. The black-eared opossum (Didelphis marsupialis 
Linnaeus) of the lowland tropics did not have any of the parasites, 
but they were quite abundant on the white-eared opossum (Didelphis 
albiventris Lund) of the subtropical and temperate zones of the 
Andes. Specimens of the common paca (Agouti paca Linnaeus) 
collected by me in the tropical zone did not have any beetles, but 
a few were found on the mountain paca (Cuniculus taczanowskii 
Stolzmann) . The greatest number of beetles, from twelve to twenty, 
were observed behind the ears of the white-eared opossum." 

Dr. Oliver Pearson had no opportunity to observe the behavior 
of the beetles that he collected; the specimens of Edrabius were 
captured from Ctenomys that had been shot, and the specimens of 
Megamblyopinus were taken from Ctenomys nests that had been 
dug out. The Amblyopinini have never before been observed in 
the nests of their hosts doubtless because the nests have not been 
examined in the past. 

Observations pertaining to the life-histories of the Amblyopinini 
have apparently not been made. Fauvel noted that the larvae of 
Edrabius philippianus had been observed on Ctenomys with the 
adults but he received no specimens. The Amblyopinini are flight- 
less, having no vestiges of wings, so are evidently closely associated 
with their hosts at almost all times. 

Host Relationships 

The species of Amblyopinini have been recorded only from 
rodents and marsupials (with the exception of one questionable bat 
association; Seevers, 1944), and it is doubtful that other mammalian 
groups harbor these parasites. The available evidence, discussed in 
more detail later, is herein interpreted as indicating that the rodents 
were the original hosts of these beetles and that their association 
with marsupials is secondary. 

The most generalized member of the group, Myotyphlus of 
Tasmania, is recorded as a parasite of "rats," probably of the house 



216 FIELDIANA: ZOOLOGY, VOLUME 37 

rat, as there are no native Tasmanian rodents. The four genera of 
the Neotropical region, occurring in subtropical and temperate 
zones, are associated with three groups of mammals: cricetine ro- 
dents, hystricomorph rodents, and marsupials. Edrabius, the South 
American genus with the most generalized morphological features, 
occurs in a limited area of Chile, western Argentina, and southern 
Peru on the burrowing hystricomorph rodent, Ctenomys. The genus 
Amblyopinus probably occurs in most subtropical and temperate 
areas of South and Central America, with the greatest number of 
its species living on cricetine rodents. Amblyopinus may be sub- 
divided into several moderately well-defined species groups. The 
jelskii and sanborni groups, with the exception of one species on 
Ctenomys, are associated with cricetine rodents; these groups are 
most abundant in the Andes of Colombia, Ecuador, and Peru, and 
occur less frequently in Central America and southern South 
America. Amblyopinus fuegensis of Tierra del Fuego, a rather dis- 
tinctive species, is a parasite of Ctenomys. Two species of the water- 
housei group occur on marsupials in the Andean region and one 
species, which I tentatively include in this group, occurs on the 
mountain paca, a hystricomorph rodent. Three species of the 
henseli group, among the most specialized members of the genus, 
are found on Brazilian marsupials. The genus Amblyopinodes, 
erected for the gahani group of species, is a highly specialized group 
associated with cricetine rodents in eastern Argentina, eastern 
Brazil, and southern Peru. The host relationships of the two species 
of Megamblyopinus are incompletely known and it is probable that 
M. mniszechi, at least, may have different hosts in different parts 
of its range. Both species of Megamblyopinus were collected from 
nests of Ctenomys in southern Peru, but inasmuch as Ctenomys does 
not occur at the type locality of M. mniszechi in central Peru some 
other rodent must serve as host in that area. 

EVOLUTIONARY CONSIDERATIONS 

There is probably too little known about the phylogenetic and 
host relationships of the Amblyopinini for us to reconstruct their 
evolutionary history, but a discussion of the topic may be profitable. 
As suggested in the previous section, the available data do not 
suggest a close parallel between the phylogeny of the beetles and 
the phyletic history of mammals. Inasmuch as only highly spe- 
cialized species of Amblyopinus are known to occur on present-day 
marsupials, it seems unlikely that the Amblyopinini began their 



SEEVERS: REVISION OF AMBLYOPININI 217 

parasitic life in association with marsupials. At some stage in the 
evolution of Amblyopinus a change in host affinities involving the 
marsupials probably occurred. 

It is difficult to reconstruct the biogeographic and phyletic 
histories of the group from known distributional and morphological 
data. The fact that the most generalized member of the tribe, 
Myotyphlus, occurs in Tasmania in association with a species of 
"rat," is in some respects confusing rather than illuminating. Per- 
haps other Old World Amblyopinini will be discovered in the future 
to throw some light on the group's history in the Old World. 

There appear to be few alternatives at present to the hypothesis 
that the Amblyopinini originated in association with primitive 
rodents, probably during the early part of the Tertiary period. 
In view of this, a brief discussion of some of the current views on 
rodent evolution is pertinent at this point. The following account 
reflects the views of Simpson (1945), Wood (1947, 1950), and Wilson 
(1951): 

The rodents arose in the Holarctic region during early Tertiary 
or very late Cretaceous times. The early rodents are generally 
placed in the suborder Sciuromorpha, a group from which the 
Myomorpha and Hystricomorpha later developed. The Hystrico- 
morpha present a number of interesting problems of evolution and 
zoogeography. This suborder has generally included a number of 
peculiar South American rodents (porcupines, guinea pigs, chin- 
chillas, pacas, agoutis, tuco-tucos, capybaras, etc.); the North 
American porcupine; the Old World porcupines of Africa, southern 
Europe, and southern Asia; and two other African families. Many 
attempts have been made to account for this zoogeographic pattern 
but without much success. Wood (1950) recently presented an 
interesting analysis of these problems and outlined his own view 
that the suborder Hystricomorpha is polyphyletic and that the 
South American genera evolved independently of the Old World 
forms. The parallelism between the Old and New World porcupines 
is very striking, and they are extremely difficult to separate on mor- 
phological grounds. Nevertheless, Wood presents a good case for the 
working hypothesis that the South American hystricomorphs evolved 
independently. Wood believed it probable that the South American 
hystricomorphs evolved from North American Paramyidae, a group 
that apparently had a Holarctic distribution in the early Tertiary. 
There is no fossil evidence whatsoever to place the hystricomorphs 
in North America so it is likely that the group differentiated in 
South America from stock that entered from the north, probably 



218 FIELDIANA: ZOOLOGY, VOLUME 37 

in late Eocene times. There is sufficient structural unity among 
the fossil hystricomorphs of the Early Oligocene to indicate that a 
single rodent type gave rise to the group. As South America was 
isolated from North America during much of the Tertiary there 
were apparently no additional rodent invasions of South America 
until the elevation of the Panamanian isthmus in Pliocene times. 
The hystricomorphs flourished during this period of isolation. The 
second migration of rodents into South America followed the ele- 
vation of the isthmus; this movement involved myomorph rodents 
of the subfamily Cricetinae. The family Cricetidae appear as fossils 
in the Oligocene of North America, Europe, and Asia; these species 
were probably the most primitive of the superfamily Muroidea. 
The Cricetinae are represented at present by the Old World hamsters 
and many New World genera that are especially prevalent in South 
America. The cricetine stock that entered South America rapidly 
became the dominant ground rodents as they were able to occupy 
ecological niches in which there was little competition. Mr. Hersh- 
kovitz informs me that a third rodent invasion has occurred in 
recent times, involving the entrance into northwestern South 
America of the sciuromorph genus Heteromys (spiny pocket-mice) 
of the family Geomyidae. 

While it is not possible to draw definite conclusions about the 
evolution of the Amblyopinini, several hypotheses may be discussed. 
There appear to be two phyletic lines within the tribe, the result 
of an early divergence. One line contains two present-day genera, 
Myotyphlus of Tasmania and Edrabius of southern South America; 
the other is comprised of Megamblyopinus, Amblyopinus, and Am- 
blyopinodes. Myotyphlus and Edrabius retain more morphological 
features of their quediine ancestors as evidenced by the structure 
of their head capsules and hind coxae. They share at least one 
specialized feature, the reduction of the eyes to one facet located 
immediately behind the antennal fossae. The Quediini have very 
large eyes that occupy much of the side of the head. The second 
phyletic line departed more from the quediine pattern and their 
head capsules, elytra, and hind coxae are more specialized; Meg- 
amblyopinus shows fewer special features than Amblyopinus and 
Amblyopinodes is the most highly specialized genus of the tribe. 

It is probable that the Amblyopinini became associated with 
rodents early in the evolutionary history of that group, perhaps in 
early Tertiary. The Myotyphlus-Edrabius stock seemingly had a 
Holarctic distribution; one may account for the occurrence of Ed- 
rabius on hystricomorph rodents in South America and Myotyphlus 







Fig. 29. a. Megamblyopinus mniszechi Solsky. b. Edrabius pearsoni sp. nov. 
c. Amblyopinus waterhonsei Fauvel. d. Myotyphlus jansoni Matthews. 



219 



220 FIELDIANA: ZOOLOGY, VOLUME 37 

in the Old World by postulating an early ancestral stock with such 
a distribution. It might seem reasonable to expect that if the pre- 
hystricomorph rodents carried these parasites into South America 
there would be more associations with present-day hystricomorphs 
than there apparently are. Edrabius may conceivably be a relict 
genus of a line of ancestral Amblyopinini that did enter South 
America with pre-hystricomorph stock; its survival may possibly be 
linked with its association with a favorable host genus, Ctenomys. 
The alternative, that Edrabius stemmed from amblyopinine stock 
that reached South America in Pliocene with the cricetine rodents, 
does not appear to be very attractive. Edrabius hardly could have 
evolved from Amblyopinus-\ike stock, so that it would be necessary 
to postulate two stocks of amblyopines on cricetine rodents and a 
subsequent transfer of one of these to the hystricomorph genus 
Ctenomys. 

The second phyletic line that which gave rise to Amblyopinus 
and related genera may well have reached South America with the 
Pliocene penetration by the cricetine rodents, although there is the 
alternative that they are derived from stock living on the native 
hystricomorphs. The first explanation requires, of course, that the 
Amblyopinini had been associated with North American rodents 
and had evolved a line somewhat different from that which gave 
rise to Edrabius. If this line was derived from stock on the hystri- 
comorphs, the ancestral type did not resemble Edrabius Very closely 
in several respects. Whether the cricetine rodents took these para- 
sites to South America with them or received them from the hystri- 
comorphs, the differentiation of Amblyopinus and Amblyopinodes 
was followed by considerable speciation accompanying that of their 
rapidly speciating hosts. There are inconsistencies in the host 
pattern, as a glance at the host list given elsewhere will show, and I 
have no explanation of these other than to postulate host transfers. 
It hardly seems plausible that ectoparasites of the nature of these 
staphylinid beetles would have adhered rigidly to host patterns 
without at times taking up new associations. Some of the species 
of Amblyopinus do not occur on cricetine rodents; several have been 
collected on Ctenomys, several on pacas, and a moderate number on 
opossums. Possibly a more complete host picture at some later 
date may provide clues to these associations. 

ACKNOWLEDGMENTS 

A substantial part of the material utilized in this revision was 
collected on the Chicago Natural History Museum Colombian 



SEEVERS: REVISION OF AMBLYOPININI 221 

Zoological Expedition (1948-52) by Mr. Philip Hershkovitz, 
Associate Curator of Mammals, and on the Peruvian Zoological 
Expedition (1953) by Mr. Celestino Kalinowski, field collector. I 
am very grateful to these men for devoting attention to collecting 
staphylinid parasites and to Mr. Hershkovitz for providing field 
notes as well as identifications of the hosts obtained on these expe- 
ditions. 

I am very much indebted to Dr. Oliver P. Pearson, Museum of 
Vertebrate Zoology, University of California, for the gift of an 
excellent collection of Amblyopinini from southern Peru. Dr. 
Pearson provided determinations of the mammalian hosts involved. 
The loan or gift of specimens by the following persons is gratefully 
acknowledged: Dr. Elli Franz, Senckenberg Museum, Frankfort 
a. M., Germany; Mr. 0. W. Steel, Maidenhead, Berkshire, England; 
and Dr. Victor van Straelen, Directeur de L/Institut Royal des 
Sciences Naturelles de Belgique. 

Subfamily Staphylininae : Tribe Amblyopinini 

Related to the tribe Quediini from which the species are separated 
on the basis of their obligate ectoparasitism. 

KEY TO GENERA OF AMBLYOPININI 

1. Eyes minute, one-faceted, located behind antennal fossae (fig. 31, a, b); gula 
and submentum as in fig. 32, a, c; hind coxae as in fig. 33, a, b 2 

Eyes multifaceted, usually large, located near posterior margin of head (fig. 
31, c, j); gula and submentum as in fig. 32, b, d, e; hind coxae as in fig. 33, 
c, d, e 3 

2. Lateral plates of ninth abdominal segment with extremely long, aciculate 
setae (fig. 44, g); South America Edrabius Fauvel 

Lateral plates of ninth abdominal segment with short setae; Tasmania. 

Myotyphlus Fauvel 

3. Sternites 3-5 (or 6) with rows of black, clavate setae (fig. 40, b); head capsule 
distinctive (figs. 30, e; 31, h); clypeus strongly deflexed in front, labrum not 
visible from above Amblyopinodes gen. nov. 

Sternites without clavate setae; head capsule otherwise, labrum visible from 
above 4 

4. Species more than 15 mm. in length; elytra longer than broad; head capsule 
relatively generalized (figs. 30, g; 31, j) Megamblyopinus gen. nov. 

Species less than 11 mm. in length; elytra broader than long; head capsule 
relatively specialized (figs. 30, c, d, f; 31, c, d, i) Amblyopinus Solsky 

Genus AMBLYOPINUS Solsky 

Amblyopinus Solsky, 1875, Hor. Soc. Ent. Ross., 11:8. 



222 FIELDIANA: ZOOLOGY, VOLUME 37 

Omaloxenus Notman, 1923, Amer. Mus. Nov., 68: 1; Costa Lima, 1936, 
Mem. Inst. Oswaldo Cruz, 31: 57 (= Amblyopinus Solsky). 

Genotype. Amblyopinus jelskii Solsky. 

Head capsule specialized (figs. 30, c, d, f; 31, c-e, i); not quediine in form. 
Eyes multifaceted, variable in size, always near posterior margin of head (figs. 
30, c; 31, c-e). Ventral surface of head distinctive (fig. 32, d); gula relatively 
short and broad; submentum relatively long; posterior tentorial pits more caudad 
in position than in the other genera (except Amblyopinodes) ; gular sutures arcuate. 
Labrum small, inconspicuous (fig. 30, c, d, /). Elytra broader than long (fig. 
33, h-j). Middle tarsi expanded and densely pilose, usually in both sexes but 
sometimes only in the male. Legs spinose. Hind coxae broad (fig. 33, d); not 
quediine in form. Lateral plates of ninth abdominal segment with numerous 
strong bristles (fig. 37, a-o). 

Remarks. This is evidently a large genus if we may judge by 
the number of new species that have been collected in recent years. 
An attempt has been made to organize species groups but this may 
be premature in view of the size of the task. Some of the hetero- 
geneity has been eliminated by the removal of two very distinctive 
species groups to new genera, Amblyopinodes and Megamblyopinus. 

Little effort has been made to construct a key to the species of 
Amblyopinus. Considerable reliance must be placed on the aedeagus 
and the terminal segments of the male abdomen for distinguishing 
closely allied species. The lateral lobes of the aedeagus are fused 
to form a single structure as in the Quediini. The fused lateral 
lobes vary considerably and are thus useful for species diagnosis. 
As the lateral lobes are difficult to illustrate from other viewpoints, 
only lateral views are given. Some of the figures show the eversible 
sac of the median lobe with its interesting structures, but this is 
not used here for taxonomic purposes because of the difficulty of 
obtaining uniform preparations for study. In some cases the ever- 
sible sac is not extruded. 

The species groups as presently organized may perhaps be sepa- 
rated by the following characteristics. The henseli group of species, 
known at present only from Brazil, is readily distinguished by the 
distinctive head capsule (figs. 30, d; 31, i) with its small, indistinctly 
faceted eyes. The other groups seem to be separable on the basis 
of the number of long setae on each half of the pronotal base: 



Fig. 30. Dorsal view of head. a. Myotyphlus jansoni Matthews, b. Edrabius 
philippianu8 Fauvel. c. Amblyopinus punae sp. no v. d. A. bequaerti Notman. 
e. Amblyopinodes gahani Fauvel. /. Amblyopinus waterhousei Fauvel. g. Megam- 
blyopinus mniszechi Solsky. 









9 




223 



224 FIELDIANA: ZOOLOGY, VOLUME 37 

One long black seta near each basal angle jelskii group 

(A. colombiae and A. spinipennis have four additional setigerous punctures 

and short, fine hairs). 
Two long black setae present A. fuegensis Arrow 

and A. hershkovitzi (this species has four setigerous punctures, but only the 

middle two have long setae). 
Three long black setae (rarely four) present A. pacae 

(one additional setigerous puncture with a fine seta is present). 
Four long black setae present A. waterhousei 

and A. kalinowskii (one, or rarely more, setigerous punctures with fine setae 

are present). 
Five or six long black setae and one or more additional setigerous punctures 

with fine setae present sanborni group 

JELSKII Group 

A group of species that occur on rodents in the Andes and the 
mountains of Central America. Distinguished from the other species 
of the genus by the presence of only one macroseta on each half of 
the pronotal base. The supraocular and infraocular setae are ar- 
ranged as in figure 31, d, except that A. schmidti, colombiae, and 
spinipennis have the arrangement (fig. 31, e) characteristic of the 
sanborni group. 

Amblyopinus jelskii Solsky 

Amblyopinus jelskii Solsky, 1875, Hor. Soc. Ent. Ross., 11: 11, pi. 1, fig. 3 
Hacienda Amable Maria, near La Merced, Junin Pro v., Peru; Matthews, 
1884, Cist. Ent., 3: 96, pi. 5, figs. 1-9; Franz, 1927, Abh. Senckenb. Nat. 
Ges., 40: 406; Costa Lima, 1936, Mem. Inst. Oswaldo Cruz, 31: 62; 
Seevers, 1944, Field Mus. Nat. Hist., Zool. Ser., 28: 161. 

Although in my earlier paper I gave a description of a species 
that I believed to be A. jelskii, I now consider it improbable that 
any of the available specimens of Amblyopinus belong to that species. 
All of the species that I consider to belong to this complex conform 
to Solsky's description but none is from a locality close to that from 
which Jelski collected the type series. I have examined one female 
specimen from the Fauvel collection determined as A. jelskii but it 
is from Callanga, Peru, the type locality of A. brandesi Kraatz, and 
is probably the specimen that Fauvel examined before synonymizing 
brandesi. There is no evidence that brandesi is conspecific with 
jelskii, and I am restoring it to the status of a valid species until 
there is evidence to the contrary. 

It is not at all unlikely that Solsky's series represented more 
than one species of Amblyopinus as several mice were involved as 












<^2 




Fig. 31. Lateral view of head. a. Myotyphlus jansoni Matthews. b.Edrabius 
philippianus Fauvel. c. Amblyopinus waterhousei Fauvel. d. A. punae sp. nov. 
e. A.sanborni Seevers. f. A. fuegensis Arrow, g. A. pacae sp. nov. h. Amblyopinodes 
gahani Fauvel. i. Amblyopinus bequaerti Notman. /. Megamblyopinus mniszechi 
Solsky. 



225 








Fig. 32. Ventral view of head. a. Myotyphlus jansoni Matthews. b.Megam- 
blyopinus mniszechi Solsky. c. Edrabius philippianus Fauvel. d. Amblyopinus 
waterhousei Fauvel. e. Amblyopinodes gahani Fauvel. 



226 



SEE VERS: REVISION OF AMBLYOPININI 227 

hosts. Jelski reported that he first encountered these beetles on 
mice that he kept in a cage at Amable Maria a species that he 
provisionally called "Mus insectivora." He also found the beetles 
in a mouse nest under a stone and on a dead mouse {"Mus lobiceps") 
in a trap. Mr. Philip Hershkovitz suggests that Oryzomys laticeps 
Lund may be one of the mice referred to by Jelski. (This species 
was reported to be one of the rodents collected by Jelski at Amable 
Maria.) 1 

Amblyopinus brandesi Kraatz. Figure 36, d, i. 

Amblyopinus brandesi Kraatz, 1900, Deuts. ent. Zeitschr., 1900:212 Callanga, 
near Cuzco, Peru; Fauvel, 1900, Revue d'Ent., 19: 64 (=jelskii Solsky). 

This species has been regarded as a synonym of A. jelskii upon 
the authority of Fauvel (1900), but there is no evidence that it is. 
The type localities of the two species are approximately three 
hundred miles apart. No host record is available for A. brandesi. 
The specimen that I presume to be Kraatz' brandesi is not conspecific 
with any of the species that I have studied. With only the female 
specimen it is difficult to characterize the species very accurately. 

Length, 8 mm. Head and pronotum reddish-brown, elytra brownish; ab- 
domen reddish-brown. Head and pronotum moderately densely and coarsely 
punctate and coarsely and closely reticulated (fig. 36, d). Eighth female sternite 
with setae as in figure 36, i. 

Material examined. One female, Callanga, Peru, from the Fauvel 
collection, Brussels Museum; determined as A. jelskii Solsky. 

Amblyopinus punae sp. nov. Figures 30, c; 31, d; 33, d; 34, a, 
b; 35, e; 36, a; 37, b. 

Length, 8 mm. Head dark reddish-brown; pronotum and abdomen paler 
reddish-brown; elytra testaceous. Head and pronotum with medium-coarse, 
relatively dense punctation. Sculpture of pronotum as in figure 36, a. Elytra 
with a dense covering of fine, flavate, recumbent setae; the apical margins with a 
few longer setae but none spinose. Eighth male sternite as in figure 35, e. Chae- 
totaxy of lateral plates of ninth male abdominal segment as in figure 37, b. Ae- 
deagus distinctive (fig. 34, a, b). 

Holotype.A male from Cailloma, Arequipa, Peru; collected 
August 25, 1939, by John M. Schmidt. Host: Akodon jelskii 
pulcherrimus Thomas. In collection of Chicago Natural History 
Museum. 

1 Oldfield Thomas, 1884. On a collection of Muridae from Central Peru. 
Proc. Zool. Soc. London, 1884: 447-458, 3 pis. 



228 FIELDIANA: ZOOLOGY, VOLUME 37 

Remarks. The specimen on which this species is based was 
determined as A. jelskii in my 1944 paper. 

Amblyopinus monticolus sp. nov. Figures 34, d; 36, g, j; 37, c. 

Distinguished from A. punae by its smaller size, paler coloration, 
pronotal sculpture, chaetotaxy of lateral plates of ninth abdominal 
segment, and aedeagus. 

Length, 5.5-6.5 mm. Coloration light-brown, with a faint reddish tinge; 
coloration uniform except that the head is a little darker in some specimens. Head 
and pronotum with fine and moderately dense punctation. Head and pronotum 
finely strigulate (fig. 36, g). Elytral setae as in A. punae. Eighth male sternite 
as in A. punae. Chaetotaxy of lateral plates of ninth male abdominal segment as 
in figure 37, c. Aedeagus as in figure 34, d. Eighth female sternite as in figure 36, j. 

Holotype. A male from Lago Suche, Moquegua, Peru; 14,600 
feet elevation; collected January 12, 1952, by Oliver P. Pearson. 
Host: From burrow of Ctenomys opimus nigriceps Thomas. In 
collection of Chicago Natural History Museum. 

Paratypes. Two specimens, same data as type. One male, 
Pampa de Titire, 29 km. northeast of Tarata, Tacna, Peru; 14,600 
feet elevation; collected April 13, 1952, by Oliver P. Pearson. Host: 
Ctenomys fulvus Philippi. In collection of Chicago Natural History 
Museum. 

Remarks. Inasmuch as the other species of the jelskii group 
are parasites of cricetine rodents, it is surprising to note that this 
species seems to be associated with Ctenomys. It is possible that 
the beetles live on rodents that inhabit Ctenomys burrows. 

Amblyopinus akodoni sp. nov. Figures 34, h; 36, b; 37, /. 

Distinguished from A. punae chiefly by its distinctive aedeagus; 
there are additional differences in sculpture of head and pronotum, 
and in the presence of semi-erect aciculate setae on the elytra. 

Length, 7 mm. Head and pronotum dark reddish-brown, abdomen lighter 
reddish-brown, elytra testaceous. Head and pronotum finely punctate. Sculpture 
of head and pronotum as in figure 36, b. Elytra rather densely clothed with 
flavate, recumbent, aciculate setae; in addition with numerous semi-erect, slightly 
stouter, aciculate setae. Tergites with a vestiture similar to that of elytra; 
their apical setae aciculate, variable in length, and not closely arranged. Lateral 
plates of ninth male abdominal segment as in figure 37, /. Aedeagus as in figure 
34, h. 

Holotype. A male from San Miguel, Tambo, Polanco, Ayacucho, 
Peru; collected November 26, 1953, by Celestino Kalinowski. Host: 
Akodon (Chroeomys) jelskii pyrrhotis Thomas (CNHM no. 75573). 
In collection of Chicago Natural History Museum. 





r ,feW >Wl 





Fig. 33. Myotyphlus jansoni Matthews: a, hind coxa; /, elytron; A;, eighth 
male sternite; I, aedeagus. Edrabius philippianus Fauvel: b, hind coxa. Megam- 
blyopinus mniszechi Solsky: c, hind coxa. Amblyopinus punae sp. nov.: d, hind 
coxa. Amblyopinodes gahani Fauvel: e, hind coxa; g, pronotum, lateral view. 
Amblyopinus kofordi sp. nov.: h, elytron. A. spinipennis sp. nov.: i, elytron. 
A. chilomysi sp. nov.: j, elytron. 



229 











Fig. 34. Aedeagi and enlarged apices of aedeagi; lateral view, a, b. Amblyo- 
pinus punae sp. nov. c. A. colombiae sp. nov. d. A. monticolus sp. nov. e, g. 
A. schmidti Seevers. /. A. kofordi sp. nov. h. A. akodoni sp. nov. 



230 



SEEVERS: REVISION OF AMBLYOPININI 231 

Amblyopinus kofordi sp. nov. Figures 33, h; 34, /; 36, c; 37, a. 

Distinguished from the other species of the jelskii group by the 
presence of semi-erect spinose setae on the elytra in addition to the 
recumbent aciculate setae, and by the distinctive aedeagus. 

Length, 7 mm. Head and pronotum light reddish-brown; elytra and abdomen 
a little paler; elytra slightly flavate. Head and pronotum with moderately dense, 
fine punctation. Sculpture of head and pronotum as in figure 36, c. Elytra 
(fig. 33, h) with a dense clothing of fine, flavate, recumbent setae and four or five 
very irregular rows of stronger, light-brown, semi-erect, spinose setae. Tergites 
with a dense clothing of moderately long, fine, recumbent hairs. Lateral plates 
of male ninth abdominal segment as in figure 37, a. Aedeagus as in figure 34, /. 

Holotype. A male from 17 miles south of Masocruz, Puno, Peru; 
13,500 feet elevation; collected May 1, 1951, by C. B. Koford. 
Host: Akodon (Bolomys) berlepschii Thomas. In collection of 
Chicago Natural History Museum. 

Paratype. A male from Pampa de Capazo, 123 km. south of 
Llave, Puno, Peru; 14,300 feet elevation; collected February 2, 
1952, by Oliver P. Pearson. Host: Akodon (Bolomys) berlepschii 
Thomas. In collection of Chicago Natural History Museum. 

Amblyopinus schmidti Seevers. Figures 34, e, g; 35, /; 36, I; 
37, d. 

Amblyopinus schmidti Seevers, 1944, Field Mus. Nat. Hist., Zool. Ser., 28: 
164; pi. 10, fig. 1; pi. 11, figs. 8-14; pi. 12, figs. 15-19, 22-23 Santa Elena, 
Chimal., Guatemala (Chicago Nat. Hist. Mus.; host, Peromyscus guate- 
malensis Merriam). 

Length, 7-8 mm. Head and pronotum reddish-brown; abdomen brownish; 
elytra light-brown. Clypeal seta absent. Head and pronotum moderately coarsely 
and densely punctate; reticulation fine-meshed but not as fine as in A. colombiae 
sp. nov. Elytra with a covering of fine, flavate, recumbent setae. Tergites with 
relatively sparse covering of short, recumbent, aciculate setae. Eighth male 
sternite with three setae on each half (fig. 35, /). Female eighth sternite with 
three (or two) setae on each half (fig. 36, I). Chaetotaxy of lateral plates of male 
ninth abdominal segment as in figure 37, d. Aedeagus as in figure 34, e, g. 

Material examined. The type series. Seven additional speci- 
mens, Finca San Rafael, 6 km. west of Mixca, Sacatepequez, Guate- 
mala, 6,900 feet elevation; collected June 28-July 1, 1948, by Rodger 
Mitchell and Luis de la Torre; from Peromyscus guatemalensis 
Merriam. 

Amblyopinus colombiae sp. nov. Figures 34, c; 36, e, m; 37, e. 

Closely related to A. schmidti but distinguished by having only 
two bristles on each half of eighth male sternite, by the arrangement 



232 FIELDIANA: ZOOLOGY, VOLUME 37 

of the setae on the female eighth sternite, and by the distinctive 
aedeagus. Both A. schmidti and A. colombiae lack the clypeal seta 
present in the other species of the jelskii group. 

Length, 8 mm. Coloration uniformly reddish-brown except for the light 
brown elytra. Clypeal seta absent. Punctation of head and pronotum fine; each 
puncture at the base of a shallow depression so that the punctures appear coarse. 
The depressions of the pronotal surface tend to be confluent so that the surface 
has numerous shallow valleys and low elevations. Sculpture of head and pronotum 
distinctive (fig. 36, e). Elytra with vestiture as in A. schmidti except that some 
setae are at posterior angles and along apical margin are longer and stronger. 
Vestiture of tergites as in A. schmidti. Chaetotaxy of lateral plates of ninth male 
abdominal segment as in figure 37, e. Eighth female sternite as in figure 36, m. 
Aedeagus as in figure 34, c. 

Holotype. A male from Las Frias, 10 km. east of Sonson, 
Antioquia, Colombia; 2,400 meters elevation; collected October 17, 
1950, by Philip Hershkovitz. Host: Thomasomys aureus nicefori 
Thomas (CNHM no. 70318). In collection of Chicago Natural 
History Museum. 
Paratypes. One male and one female, same data as type. 

Amblyopinus piurae sp. nov. Figure 35, g. 

Most closely allied to A. colombiae; distinguished by its more 
slender build, paler coloration, more coarsely meshed reticulation 
of pronotum and minor differences in aedeagus. 

Length, 5.5-6 mm.; pronotal width, 2 mm. Coloration pale rufo-flavate. 
Clypeal seta, if present, very minute. Pronotal punctation moderately coarse 
and dense; reticulation medium-meshed. Elytra and tergites with fine, moderately 
dense, aciculate, recumbent setae. Eighth male sternite with two dark setae on 
each side of emargination. Aedeagus as in figure 35, g. 

Holotype. A male from Tambo, Huancabamba, Piura, Peru; 
2,880 meters elevation; collected May 8, 1954, by Celestino Kali- 
nowski. Host: Thomasomys sp. (CNHM no. 81334). In collection 
of Chicago Natural History Museum. 

Paratype. One male, same data as type. 

Amblyopinus ancashi sp. nov. Figure 35, h. 

Distinguished from the other species of the jelskii group by its 
distinctive aedeagus. 

Length, 6.5-7 mm. Coloration of head and pronotum reddish-brown; abdomen 
pale brown; elytra flavo-testaceous. Head and pronotum with a medium-meshed 
reticulation. Elytra with a relatively dense vestiture of coarsely aciculate, 
recumbent setae; many of the setae with the same color as the elytra but some 
light brown. Tergites with a medium-dense covering of recumbent aciculate 






Fig. 35. Aedeagi, enlarged apices of aedeagi, and eighth male sternites. 
a, b. Amblyopinus hershkovitzi sp. nov. c. A.fuegensis Arrow, d. A. spinipennis sp. 
nov. e. A. punae sp. nov. /. A. schmidti Seevers. g. A. piurae sp. nov. 
h. A. ancashi sp. nov. 



233 



234 FIELDIANA: ZOOLOGY, VOLUME 37 

setae, most of which are a little paler than the tergites. Apical setae of elytra 
and tergites longer than diskal setae for the most part. Eighth male sternite 
with one dark seta on each side of emargination. Aedeagus as in figure 35, h. 

Holotype. A male from Quilcayhanca, Huaras, Ancash, Peru; 
4,000 meters elevation; collected February 20, 1954, by Celestino 
Kalinowski. Host: Phyllotis darwini Waterhouse (CNHM no. 
81219). In collection of Chicago Natural History Museum. 

Paratypes. Two specimens, same data as type. 

Amblyopinus sp. 

A female specimen of the jelskii group, related to the foregoing 
species but distinct from it, was collected near Puno, Puno, Peru, 
by Oliver P. Pearson, December 19, 1951, from Phyllotis (Aulis- 
comys) sublimis Thomas. 

Amblyopinus spinipennis sp. nov. Figures 33, i; 35, d; 37, g. 

Distinguished from the other species by a row of semi-erect 
spinose setae on the apical margins of the elytra and by its aedeagus. 

Length, 7 mm. Head, pronotum, and abdomen reddish-brown; elytra tes- 
taceous. Pronotum with the usual black seta of the jelskii group near each basal 
angle but with four additional setigerous punctures along basal margin, each bear- 
ing a short fine seta. Elytra clothed with fine, flavate, recumbent setae and with 
an apical row of moderately strong, semi-erect, spinose setae (fig. 33, i). Tergites 
with recumbent aciculate setae that are more dense laterally; apical margins of 
tergites with moderately stout, aciculate setae. Chaetotaxy of lateral plates of 
ninth male abdominal segment as in figure 37, g. Aedeagus as in figure 35, d. 

Holotype. A male from Marcapata, Limapunco, Cuzco, Peru; 
2,400 meters elevation; collected July 1, 1953, by Celestino Kali- 
nowski. Host: Akodon aerosus Thomas (CNHM no. 75470). 

Amblyopinus longus Franz 

Amblyopinus longus Franz, 1930, Senckenbergiana, 12: 74, figs. 6, 7 Hansa, 
Santa Catharina, Brazil (Senckenberg Mus.; from Oxymycterus rufus 
Desmaret); Fonseca, 1939, Mem. Inst. Butantan, 12: 191. 

This species, which evidently belongs to the jelskii group, 
is distinguished by the chaetotaxy of the tergites and the eighth 
male sternite. Most of the tergites have four dark setae on each 
half of their apical margins; the other species of the group have at 
most two such setae and the outer one is frequently fine and incon- 
spicuous. The eighth male sternite has seven or eight dark setae 
on each half; the other species have one or two as a rule, except 
that A. schmidti has three. The aedeagus is probably distinctive 
but I have had no opportunity to study it. 













f I 





Fig. 36. Pronotal sculpture (a-h); eighth female sternites (i-m). a. Am- 
blyopinus punae sp. nov. b. A. akodoni sp. nov. c. A. kofordi sp. nov. d, i. 
A. brandesi Kraatz. e, m. A. colombiae sp. nov. /. A. fuegensis Arrow, g, j. 
A. monticolus sp. nov. h, k. A. hershkovitzi sp. nov. I. A. schmidti Seevers. 






235 




Fig. 37. Ventral views of lateral plates of ninth male abdominal segments. 
a. Amblyopinus kofordi sp. nov. b. A. punae sp. nov. c. A. monticolns sp. nov. 
d. A. schmidti Seevers. e. A. colombiae sp. nov. /. A. akodoni sp. nov. 
g. A. spinipennis sp. nov. h. A. hershkovitzi sp. nov. i. A. waterhousei Fauvel. 
j. A. montivagans sp. nov. k. A. sanborni Seevers. I. A. chilomysi sp. nov. 
to. A. emarginatus sp. nov. n. A. huilae sp. nov. o. A. oryzomysi sp. nov. 



236 



SEEVERS: REVISION OF AMBLYOPININI 237 

Length, 6 mm. Coloration rufo-flavate; a relatively pale species. Clypeal 
seta present. Setae in vicinity of eyes as in A. punae. Head and pronotum with 
fine medium-meshed reticulation. Pronotal base with one black seta near basal 
angle and a much finer seta more medially placed. Elytra and tergites with pale, 
recumbent, aciculate setae. Tergites with dark setae on each half of apical margins 
as follows: tergite 2 with one; tergites 3-7 with four; tergite 8 with two on apical 
margin and three along lateral margin. Eighth male sternite with three black 
setae near apical margin and four or five near lateral margin. 

Material examined. One male paratype, Hansa, Santa Catha- 
rina, Brazil. 

Remarks. Fonseca (1939) recorded this species from Sao Paulo, 
Brazil, with Akodon (Thaptomys) nigrita Lichtenstein. It is possible 
that this is a record of a species closely related to A. longus. 

FUEGENSIS Group 

Amblyopinus fuegensis Arrow occupies a somewhat isolated 
position in the genus and is probably not very closely linked to the 
foregoing group. The locations of the eyes and the two near-by 
setae are different (fig. 31, /) ; the pronotal base has two black setae 
on each half; the elytra are relatively long for the genus; the eighth 
male sternite has a number of long dark setae on each half instead 
of the usual one to three setae. 

At present only one species is placed in this group but there 
may well be others in southern South America. 

Amblyopinus fuegensis Arrow. Figures 31, /; 35, c; 36, /. 

Amblyopinus fuegensis Arrow, 1907, Ann. Mag. Nat. Hist., (7), 19: 126 

Useless Bay, Tierra del Fuego, Chile; Seevers, 1944, Field Mus. Nat. Hist., 

Zool. Ser., 28: 163, pi. 12, fig. 21. 

Length, 9 mm. Light-brown throughout, with only a faint reddish tinge. 

Head with relatively coarse and dense punctation; pronotum with finer and 

sparser punctures. Head and pronotal reticulation obsolescent (fig. 36,/). Elytra 

with a clothing of fine, flavate, aciculate setae. Tergites relatively sparsely clothed 

with medium-long, recumbent, aciculate setae. Aedeagus distinctive (fig. 35, c). 

Material examined. One male, Riesco Island, Magallanes, Chile. 
Host: Ctenomys magellanicus dicki Osgood. 



HERSHKOVITZI Group 

A. hershkovitzi is such a distinctive species that it is placed in a 
separate group. The characteristics as given in the description 
below should serve to identify this species. 



238 FIELDIANA: ZOOLOGY, VOLUME 37 

Amblyopinus hershkovitzi sp. nov. Figures 35, a, b; 36, h, k; 
37, h. 

Length, 7.5-9 mm. Head and pronotum light-brown, with only a slight 
reddish tinge; abdomen medium reddish-brown; elytra brown, only slightly paler 
than the other parts. Punctation of head and pronotum fine and relatively sparse. 
Sculpture of head and pronotum fine and close (fig. 36, h). Pronotal base with 
four setigerous punctures on each half; the middle two punctures bear long black 
setae, the others have short, fine hairs. Eyes relatively small, reduced to six or 
seven indistinct facets. Occipital line absent. 

Elytra clothed with recumbent, yellowish-brown hairs (stiffer than in most 
species), and stouter brown, weakly spinose setae. Tergites clothed with moder- 
ately long, aciculate setae, more dense laterally than in the middle; tergites 
without semi-erect setae. Apical margins of tergites with relatively sparse acic- 
ulate setae of two lengths. Eighth male sternite with two setae (sometimes 
only one) on each half. Chaetotaxy of lateral plates of ninth male segment as in 
figure 37, h. Aedeagus as in figure 35, a, b. Female eighth sternite with only one 
seta on each half (fig. 36, k). 

Holotype. A male from Valdivia, Antioquia, Colombia; 1,400 
meters elevation; collected June 14, 1950, by Philip Hershkovitz. 
Host: Heteromys anomalus Thompson (CNHM nos. 70474-76). In 
collection of Chicago Natural History Museum. 

Paratypes. Two specimens, same data as type. 



SANBORNI Group 

A group of species that occur on cricetine rodents in the Andes. 
The species are probably most easily distinguished by the presence 
of five or six long black setae on each half of the pronotal base. 
In the species of the sanborni group the posterior surface of the head 
is more deeply concave than in the species of the jelskii group, but 
this is not an easy character to use for diagnostic purposes. The 
infraocular seta of the head (fig. 31, e) is located in an angulate 
projection of the head but this condition occurs in several species 
of the jelskii group (schmidti, colombiae, and spinipennis) as well 
as in A. hershkovitzi. 

Amblyopinus sanborni Seevers. Figures 31, e; 37, k; 38, c, h. 

Amblyopinus sanborni Seevers, 1944, Field Mus. Nat. Hist., Zool. Ser., 28: 

166 Segrario, Puno, Peru (Chicago Nat. Hist. Mus.; from a bat, Carollia 

perspicillata Linnaeus). 

Length, 5.5-7 mm. Head and pronotum moderately reddish-brown; elytra 

and abdomen paler brown. Head and pronotum with coarse, "crateriform" 

punctures and a close-meshed reticulation. Pronotal base with six or seven 

setigerous punctures on each half and with five or six long black setae and one 



SEEVERS: REVISION OF AMBLYOPININI 239 

or more short fine hairs. Elytra with fine to moderately coarse aciculate setae. 
Tergites with a moderately uniform clothing of fine aciculate setae (semi-recum- 
bent), short to medium long. Pronotal base shallowly emarginate; surface feebly 
depressed in front of emargination. Eighth male sternite with two black setae 
on each half (fig. 38, h). Lateral plates of ninth male abdominal segment as in 
figure 37, k. Aedeagus as in figure 38, c. 

Material examined. The type. Four specimens from Marcapata, 
Limapunco, Cuzco, Peru; 2,400 meters elevation; collected July 13, 
1953, by Celestino Kalinowski; from Thomasomys sp. (CNHM no. 
75369). Seven specimens, Hacienda Cadena, Marcapata, Cuzco, 
Peru; collected September 1, 1950, by Celestino Kalinowski; from 
Oryzomys keaysi Allen (CNHM no. 68628). 

Remarks. This species was originally based on one female 
specimen collected from a bat, Carollia perspicillata. This host 
record was so unusual that it was suspected of being a chance 
association. The suspicion seems to have been justified, for this 
species has since been taken from species of two genera of mice, 
Thomasomys and Oryzomys. The specimens on the latter mouse 
are smaller (5.5-6.5 mm.) than those from Thomasomys (7 mm.), 
but I can note no other basis for separating the specimens. 

Amblyopinus emarginatus sp. nov. Figures 37, m; 38, /. 

Distinguished from A. sanborni by having the pronotal base 
emarginate and by its distinctive aedeagus. 

Length, 7-7.7 mm. Head and pronotum moderately dark reddish-brown; 
the elytra and abdomen light brown. Head and pronotum with dense, irregularly 
arranged, crateriform punctation; surface reticulated. Chaetotaxy of pronotal 
base as in A. sanborni. Elytra with a uniform covering of fine flavate setae, 
longer on sides and near posterior angles. Tergites with vestiture similar to that 
of elytra; apical row of setae not distinctive. Pronotal base broadly and moder- 
ately deeply emarginate; surface in front of emargination broadly depressed. 
Eighth male sternite with two setae. Lateral plates of ninth male segment as in 
figure 37, m. Aedeagus as in figure 38, /. 

Holotype. A male from San Antonio, San Agustin, Huila, 
Colombia; collected during August-October, 1951, by Philip Hersh- 
kovitz. Host: Thomasomys laniger Thomas. In collection of 
Chicago Natural History Museum. 

Paratype. One male, same data as type. 

Amblyopinus chilomysi sp. nov. Figures 33, /; 37, I; 38, a, b. 

This and the three following species are distinguished from A. 
sanborni and A. emarginatus by the presence of semi-erect spinose 
setae on the elytra in addition to the usual recumbent, aciculate 



240 FIELDIANA: ZOOLOGY, VOLUME 37 

type. A. chilomysi is also distinguished from the two preceding 
species by the non-crateriform punctures and the strigulate sculpture 
of the head. The aedeagus and the chaetotaxy of the ninth lateral 
plates of the male are distinctive. 

Length, 7 mm. Coloration light brown, with a faint reddish tinge. Head 
with coarse, moderately dense punctation. Sculpture of head consisting of wavy 
lines with few cross connections. Pronotum with moderately coarse punctures 
accompanied by confluent depressions; the surface thus with alternating valleys 
and ridges. Elytra with fine, flavate, aciculate setae and scattered, pale brown, 
weakly spinose setae, semi-erect but not conspicuously different from the recum- 
bent setae (fig. 33, j). Tergites with recumbent, aciculate setae; their apical mar- 
gins with a row of relatively long, aciculate setae some of which are long, others 
intermediate in length. Chaetotaxy of lateral plates of ninth male abdominal 
segment as in figure 37, I. Aedeagus distinctive (fig. 38, a, b). 

Holotype. A male from Rio Balcones, Guasca, Cundinamarca, 
Colombia; 2,750 meters elevation; collected June 4, 1952, by Philip 
Hershkovitz. Host Chilomys sp. (CNHM no. 71636). 

Amblyopinus montivagans sp. nov. Figures 37, j; 38, e. 

Close to A. chilomysi; distinguished by its aedeagus as well as 
by several minor differences. 

Length, 7.5 mm. Coloration light brown; head and pronotum with slight 
reddish tinge. Head with coarse, moderately dense, non-crateriform punctation. 
Pronotum finely and moderately densely punctate; surface without depressions 
so it is not undulating as in A. chilomysi. Head and pronotum with fine, wavy 
lines having few cross connections. Elytra with aciculate setae and semi-erect, 
spinose setae as in A. chilomysi. Tergites with small and rather fine, aciculate 
setae and stouter, darker, longer setae. Pronotal base with five, long, black setae 
and one, short, fine seta on each half. Pronotal base scarcely emarginate. Form 
of elytra more as in figure 33, h than as in A. chilomysi (fig. 33, j). Chaetotaxy 
of lateral plates of ninth male abdominal segment as in figure 37, j. Aedeagus 
as in figure 38, e. 

Holotype. A male from San Cristobal, Bogota, Cundinamarca, 
Colombia; 2,800 meters elevation; collected July 5, 1952, by Philip 
Hershkovitz. Host: Chilomys sp. (CNHM no. 71630). 

Amblyopinus oryzomysi sp. nov. Figures 37, o; 38, g, i. 

Distinguished from the two preceding species by the reticulate 
sculpture of head and pronotum, by the row of spinose setae on the 
apical margin of the tergites, and by the aedeagus. 

Length, 7-8 mm. Head, pronotum, and abdomen reddish-brown; elytra 
brown. Head and pronotum densely punctate; the punctures coarse and crateri- 
form. Head and pronotum with a medium-coarse reticulation. Elytra densely 














Fig. 38. Aedeagi, enlarged apices of aedeagi, and eighth male sternites. 
a, b. Amblyopinus chilomysi sp. no v. c,h.A. sanborni Seevers. d. A. huilae sp. nov. 
e. A. montivagans sp. nov. /. A. emarginatus sp. nov. g, i. A. oryzomysi sp. nov. 



241 






242 FIELDIANA: ZOOLOGY, VOLUME 37 

clothed with unusually short, aciculate setae and with moderately numerous 
short, light-brown, semi-erect, spinose setae. Tergites with a sparse covering of 
relatively short, recumbent, aciculate setae; their apical margins with a continuous 
row of short spinose setae (comb-like, but scarcely a ctenidium). Chaetotaxy of 
the lateral plates of ninth male abdominal segment as in figure 37, o. Aedeagus 
as in figure 38, g, i. 

Holotype. A male from Las Frias, 10 km. east of Sonson, 
Antioquia, Colombia; 2,400 meters elevation; collected October 17, 
1950, by Philip Hershkovitz. Host: Oryzomys albigularis Tomes 
(CNHM no. 70456). In collection of Chicago Natural History 
Museum. 

Paratypes. Two specimens, same data as type. Two specimens 
from Guapantal, Urrao, Antioquia, Colombia; 2,200 meters eleva- 
tion; collected April 13, 1951, by Philip Hershkovitz; from Oryzomys 
albigularis Tomes. One specimen from San Antonio, San Agustin, 
Huila, Colombia; 2,200 meters elevation; collected August 27, 1951, 
by Philip Hershkovitz; from Oryzomys albigularis Tomes. 

Amblyopinus huilae sp. nov. Figures 37, n; 38, d. 

Distinguished from the other species of the sanborni group by its 
distinctive aedeagus and by having two or three long setae (four 
setigerous punctures) on each half of the pronotal base. 

Length, 6.3 mm. Head, pronotum, and elytra testaceous; abdomen slightly 
rufo-testaceous. Head with dense, coarse, non-crateriform punctation, and with 
a coarse-meshed reticulation. Pronotum with medium-coarse, medium-dense 
punctures, and with fine, inconspicuous reticulation. Pronotal base with four 
setigerous punctures on each half and only two or three long black setae. Elytral 
setae as in A. montivagans. Tergites with rather uniform covering of moderately 
long, fine, aciculate setae, their apical rows of spinose setae nearly as in A. ory- 
zomysi. Chaetotaxy of the lateral plates of ninth male segment as in figure 37, n. 
Aedeagus as in figure 38, d. 

Holotype. A male from San Antonio, San Agustin, Huila, 
Colombia; 2,250 meters elevation; collected September 4, 1951, by 
Philip Hershkovitz. Host: Chilomys sp. (CNHM no. 71494). In 
collection of Chicago Natural History Museum. 



WATERHOUSEI Group 

The species of this group are not markedly different from those 
of the sanborni group but do exhibit some distinctive qualities: the 
posterior surface of the head is deeply concave lateral to the neck; 
the eyes and the two closely associated setae are arranged as in 
figure 31, c; the pronotal base has three or four long black setae 



SEEVERS: REVISION OF AMBLYOPININI 243 

on each half; the elytra and tergites are very sparsely setose; the 
species are the largest of the genus. 

The three species assigned to this group occur in the Andes. 
A. waterhousei and A. kalinowskii are associated with the white- 
eared opossum, while A. pacae occurs on the mountain paca. A. 
pacae may be incorrectly placed but this cannot be decided until 
more material of this genus is available. 

Amblyopinus waterhousei Fauvel. Figures 29, c; 30, /; 31, c; 
32, d; 37, i; 39, a-c. 

Amblyopinus waterhousei Fauvel, 1900, Revue d'Ent., 19: 64 Riobamba, 
Chimborazo, and Cuenca, Azuay, Ecuador (Brit. Mus.; no host given); 
Costa Lima, 1936, Mem. Inst. Oswaldo Cruz, 31:63; Seevers, 1944, 
Field Mus. Nat. Hist., Zool. Ser., 28: 165, pi. 10, figs. 4, 5. 
Length, 8.2-10.5 mm. Head reddish-brown; remainder of body light brown 
with a slight rufous or flavous tinge. Head punctures relatively coarse and dense; 
those of pronotum appreciably finer and a little sparser. Sculpture of head and 
pronotum a very fine, parallel strigulation with few cross-connections (similar to 
that of A. hershkovitzi, fig. 36, h). Pronotal base with four long, black setae on 
each half and one, or rarely more, setigerous punctures bearing fine setae; one or 
more of the long setae may be reduced. Elytral punctation unusually coarse. 
Elytra with fine, relatively sparse, aciculate setae arising from tubercles; the sur- 
face as a result is unusually rough. Tergites with a sparse vestiture of short, 
fine hairs. Eighth male sternite with one seta on each half (fig. 39, c). Lateral 
plates of ninth male abdominal segment as in figure 37, i. Aedeagus as in figure 
39, a, b. 

Material examined. One female from the Fauvel collection, 
Brussels Museum, labeled "Cuenca, Equateur." Seventeen speci- 
mens, San Antonio, San Agustin, Huila, Colombia; 2,200 meters 
elevation; collected September 10, 1951, by Philip Hershkovitz. 
Five specimens, El Calvario, Meta, Colombia; collected November 
20, 1939, by Ernesto Osorno (Mus. Comp. Zool. and Chicago Nat. 
Hist. Mus. collections). Two specimens, Bogota, Colombia; col- 
lected by Philip Hershkovitz. Host: Didelphis albiventris Lund, 
the white-eared opossum (Didelphis paraguayensis "Oken" and D. 
azarae of authors, not Temminck). 

Amblyopinus kalinowskii sp. nov. Figure 39, d. 

Closely related to A. waterhousei; distinguished by the structure 

Jof its aedeagus (fig. 39, d). 
Length, 9-10 mm. Coloration reddish-brown throughout; head a little darker. 
Holotype. A male from Marcapata, Limapunco, Cuzco, Peru; 
2,400 meters elevation; collected July 15, 1953, by Celestino Kali- 



244 FIELDIANA: ZOOLOGY, VOLUME 37 

nowski. Host: Didelphis albiventris Lund (CNHM no. 75103). 
In collection of Chicago Natural History Museum. 

Paratypes. Six specimens, same data as type, except that one 
specimen was taken from a second opossum on July 16, 1953. 

Amblyopinus pacae sp. nov. Figures 31, g; 39, e, f. 

Tentatively assigned to the waterhousei group, this species is 
distinguished by its aedeagus, the chaetotaxy of the eighth male 
sternite, and the presence of only three, long, black setae on each 
half of the pronotal base. 

Length, 9 mm. Coloration reddish-brown throughout. Arrangement of the 
setae on the side of the head as in figure 31, g; the structure of the head capsule 
very much like that of A. waterhousei. Pronotal base with three long setae 
(rarely four) and one short seta on each half. Elytra sparsely clothed with fine, 
flavate, aciculate setae, longer near outer posterior angles and on lateral margins. 
Tergites with an unusually sparse vestiture of short, fine hairs. Eighth male 
sternite as in figure 39, e. Aedeagus as in figure 39, /. 

Holotype. A male from Marcapata, Cuzco, Peru; 3,100 meters 
elevation; collected August 2, 1953, by Celestino Kalinowski. Host: 
Cuniculus taczanowskii Stolzmann, the mountain paca (CNHM no. 
75125). In collection of Chicago Natural History Museum. 

Paratypes. Four specimens, same data as type. 

Amblyopinus sp. 

A female specimen from Rio Balcones, Guasca, Colombia, col- 
lected on June 16, 1952, by Philip Hershkovitz from Cuniculus 
taczanowskii, is closely related to the preceding species. 



HENSELI Group 

The three species of this group occur on Brazilian marsupials. 
It is difficult to decide whether or not these species arose from the 
same stock within Amblyopinus as the species of the waterhousei 
group did. The head capsule of henseli and related species has 
undergone greater specialization in some respects (fig. 30, d; 31, i). 
The eyes are quite small and indistinctly faceted. 

For descriptions of bequaerti and marmosae see Seevers (1944). 



Fig. 39. Aedeagi, enlarged apices of aedeagi, and eighth male sternite. a-c. 
Amblyopinus waterhousei Fauvel. d. A. kalinowskii sp. nov. e,f. A. pacae sp. nov. 









245 








mm 




'il'i'ii'lii'ilVi'i' 1 ','/ 1 , 1 

mm 




/Hi! Vi 11 !' I/, 

iViVifiJi'i'Ji'ili 



Ifil'if'/WI 







I)! y 



'jl'iui l"[ 




" ''ii'i/f/ii 

" '/'IVMI/ 



ffr/f/ 




I' (If 'If' I In 

'V '.hi 



i 'r, 





Fig. 40. Amblyopinodes danger Franz: a, sternites 3-7; d, second and third 
tergites; i, eighth male sternite; I, basal angle of pronotum. A. gahani Fauvel: 
c, second and third tergites; e, sternites 5-7; h, eighth male sternite; k, basal 
angle of pronotum. A. piceus Brethes: b, sternites 3-6; /, eighth male sternite; 
m, basal angle of pronotum. A. peruvianus sp. nov.: g, eighth male sternite. 
A. similis sp. nov.: j, eighth male sternite. 



246 



SEEVERS: REVISION OF AMBLYOPININI 247 

Amblyopinus henseli Kolbe 

Amblyopinus henseli Kolbe, 1911, Deuts. ent. Nat. Bibl., 2: 117 Rio Grande 
do Sul or Rio de Janeiro, Brazil (Berlin Mus.; no host); Franz, 1927, 
Abh. Senckenb. Naturf. Ges., 40: 405, 4 figs.; Costa Lima, 1936, Mem. 
Inst. Oswaldo Cruz, 31: 64; Zikan, 1939, Rev. Ent., 10: 222, 8 figs.; 
Fonseca, 1939, Mem. Inst. Butantan, 12: 191; Seevers, 1944, Field Mus. 
Nat. Hist., Zool. Ser., 28: 167. 

Records. Serra dos Orgaos, Brazil (Franz); Angra dos Reis, 
Brazil, and Tijuca, Rio de Janeiro, Brazil (Costa Lima) ; Sao Paulo, 
Brazil (Fonseca). 

Host records. Metachirus opossum Linnaeus (Franz, Zikan). 
Didelphis marsupialis Linnaeus (Costa Lima, Fonseca). 

Amblyopinus bequaerti Notman. Figures 30, d; 31, i. 

Omaloxenus bequaerti Notman, 1923, Amer. Mus. Nov., no. 68 Itatiaya, 
Rio de Janeiro, Brazil (Amer. Mus. Nat. Hist.; Monodelphis sp.). 

Amblyopinus bequaerti Notman; Costa Lima, 1936, Mem. Inst. Oswaldo 
Cruz, 31: 64 (=henseli Kolbe); Seevers, 1944, Field Mus. Nat. Hist., 
Zool. Ser., 28: 168, pi. 10, fig. 6, pi. 12, fig. 24 (A. bequaerti a valid species). 

Material examined. A male paratype. 
Amblyopinus marmosae Seevers 

Amblyopinus marmosae Seevers, 1944, Field Mus. Nat. Hist., Zool. Ser., 28 : 
169, pi. 12, figs. 25, 26 Apiahy, Sao Paulo, Brazil (Mus. Comp. Zool.; 
Marmosa incana paulensis Tate). 



Specimens of the following species have not been available for 
study and I have not been able to determine its position from the 
original description. 

Amblyopinus angustus Arrow 

Amblyopinus angustus Arrow, 1907, Ann. Mag. Nat. Hist., (7), 19: 125 
Monte Roraima, British Guiana (Brit. Mus.; host not recorded). 

Genus AMBLYOPINODES gen. nov. 

Genotype. Amblyopinodes gahani Fauvel. 

Distinguished from the other genera by its distinctive head 
capsule (figs. 30, e; 31, h; 32, e) and by the presence of rows of black, 
claviform setae on three or four sternites (fig. 40, a, b) . The clypeus 
is strongly deflexed and the labrum invisible from above. 



248 FIELDIANA: ZOOLOGY, VOLUME 37 

Amblyopinodes evidently had its origin in common with Am- 
blyopinus. During its evolution the head capsule acquired a form 
surprisingly like that of certain other parasitic beetles, the beaver 
parasite {Platypsyllus castoris Ritsema) in particular. The presence 
of black, clavate "setae" on the under side of the abdomen is a 
very interesting feature of this genus. These structures are evidently 
movable at the base and they can scarcely be called a ctenidium. 
Their function is unknown. 

Species of the genus are now known to occur in eastern Argentina, 
southern Brazil, and southern Peru, in all cases on cricetine rodents. 

Amblyopinodes gahani Fauvel comb. nov. Figures 30, e; 31, h; 
32, e; 40, c, e, h, k; 41, a. 

Amblyopinns gahani Fauvel, 1901, Revue d'Ent., 20: 6 La Plata, Argentina 
(Brit. Mus.). 

This species probably has a narrower range than students of 
the group believed. It is almost certain that few of the specimens 
identified as A. gahani in the past have actually belonged to that 
species. Costa Lima (1936) placed A. claviger Franz and A. piceus 
Brethes in synonymy with A. gahani but I do not believe that he 
was justified in doing so. I have studied series of specimens from 
the type localities of both claviger and piceus and have compared 
them with a syntype of gahani. There is little doubt that three 
species are present and I am reasonably certain that the three 
named species are valid. 

A. gahani is distinguished from claviger, to which it seems to be 
most closely related, by its aedeagus (fig. 41, a) and by certain 
features of the abdominal chaetotaxy (fig. 40, c, e, h). 

Material examined. One male syntype from the Fauvel col- 
lection, Brussels Museum. 

Amblyopinodes piceus Brethes comb. nov. Figures 40, b, f, m; 
41, c. 

Omaloxenus piceus Brethes, 1926, An. Mus. Nac. Hist. Nat. Bernardino Riva- 

davia, 34: 17 Misiones, Argentina ("sur des rats"). 
Amblyopinus piceus Brethes, Costa Lima, 1936, Mem. Inst. Oswaldo Cruz, 

31: 59 (=gahani Fauvel); Fonseca, 1939, Mem. Inst. Butantan, 12: 

191 (det. as gahani Fauvel). 

This species has generally been confused with A. gahani, al- 
though there are several good distinguishing features. The number 
and arrangement of the black clavate setae on the sternites are 



SEEVERS: REVISION OF AMBLYOPININI 249 

distinctive features (fig. 40, 6). The aedeagus (fig. 41, c) is distinc- 
tive, and the base of the pronotum has four long setae (fig. 40, m) 
as contrasted with three in gahani and claviger. 

This species is a parasite on the water rat, Nectomys squamipes. 

Records. Therezopolis, Brazil, from Nectomys squamipes Brants 
(Costa Lima). Sao Paulo, Brazil, from Nectomys squamipes 
Brants (Fonseca). 

Material examined. Seven specimens, Caraguatay, Misiones, 
Argentina, collected September 10, 1926, by Marshall Field Brazilian 
Expedition of Chicago Natural History Museum; from Nectomys 
squamipes Brants. 

Amblyopinodes travassosi Costa Lima comb. nov. 

Amblyopinus travassosi Costa Lima, 1927, C. R. soc. Biol., 79: 842 Japuhyba, 
Rio Janeiro, Brazil (Inst. Oswaldo Cruz; "rato do matto"); Costa Lima, 
1936, Mem. Inst. Oswaldo Cruz, 31: 61, text fig. 2; Fonseca, 1939, Mem. 
Inst. Butantan, 12: 191. 

Costa Lima distinguished this species from gahani (actually in 
all probability from claviger and piceus) by its smaller size (4-5.5 
mm. in length) and the structure of its aedeagus. I have not seen 
specimens of this species. 

Records. Costa Lima recorded this species from Santa Catha- 
rina, Brazil, from a "rato grande castanho," and from Sao Paulo, 
Brazil, from a "rato selvagem." Fonseca recorded A. travassosi 
from the state of Sao Paulo from Thaptomys nigrita Lichtenstein, 
and from Oxymycterus judex Thomas. 

Amblyopinodes claviger Franz comb. nov. Figures 40, a, d, i, I; 

41, b. 

Amblyopinus claviger Franz, 1930, Senckenbergiana, 12: 71, text figs. 1-5 
Hansa, Santa Catharina, Brazil (Senckenberg Mus.; from Oxymycterus 
rufus Desmaret); Costa Lima, 1936, Mem. Inst. Oswaldo Cruz, 31: 59 
(= gahani Fauvel). 

Although this species is doubtless close to A. gahani I believe 
that the differences justify recognizing it as a distinct species. There 
are differences in chaetotaxy (compare fig. 40, c and d, the second 
and third tergites; and fig. 40, a and e, the fifth to seventh sternites), 
and in the structure of the aedeagus (compare fig. 41, a and b). 

Material examined. Two paratypes, Hansa, Santa Catharina, 
Brazil. Seven specimens, Nova Teutonia, Santa Catharina, Brazil, 
collected by Fritz Plaumann, no host given. 



250 



FIELDIANA: ZOOLOGY, VOLUME 37 








Fig. 41. Aedeagi and apices of aedeagi. 
b. A. danger Franz, c. A. piceus Brethes. 
A. similis sp. nov. 



a. Amblyopinodes gahani Fauvel. 
d. A. peruvianus sp. nov. e, f. 



Amblyopinodes peruvianus sp. nov. Figures 40, g; 41, d. 

Distinguished from the foregoing species of the genus by having 
a lesser number of black clavate setae on the sternites and by the 
structure of its aedeagus. 

Length, 5.25-6 mm. Pronotal punctation medium coarse and moderately 
dense. Pronotal reticulation obsolescent. Pronotal base with four long setae 
on each half (the third in the series is occasionally reduced in length). Second 



SEEVERS: REVISION OF AMBLYOPININI 251 

tergite with three or four long setae on each half of its apical margin; tergites 
3-8 with three setae on each half; tergites 6 and 7 with one seta near each lateral 
margin ; tergite 8 with a pair of setae near lateral margin about midway back and 
one sub-basal seta. Third sternite with six or eight black clavate setae, fourth 
sternite with four or five, and fifth sternite with one such seta. Aedeagus as in 
figure 41, d. Eighth male sternite as in figure 40, g. 

Holotype. A male from Quince Mil, Cuzco, Peru; 680 meters 
elevation; collected August 10, 1953, by Celestino Kalinowski. 
Host: Oryzomys sp. (CNHM no. 75272). In collection of Chicago 
Natural History Museum. 

Paratypes. Six specimens, same data as type, and one specimen 
from the same locality, collected on August 8, 1953, on another 
of Oryzomys sp. (CNHM no. 75270). 

Amblyopinodes similis sp. nov. Figures 40, j; 41, e, f. 

Very similar to A. peruvianus; distinguished from it by the structure of its 
aedeagus (fig. 41, e, /) and the eighth male sternite (fig. 40, j). 

Holotype. A male from Hacienda Cadena, Marcapata, Cuzco, 
Peru; 1,000 meters elevation; collected January 12, 1949, by Celes- 
tino Kalinowski, from Oxymycterus juliacae Allen (CNHM no. 
65704). In collection of Chicago Natural History Museum. 

Paratype. One specimen, same data as type. 

Genus MEGAMBLYOPINUS gen. nov. 

Genotype. Megamblyopinus mniszechi Solsky. 

Megamblyopinus evidently stemmed from the same evolutionary 
line that gave rise to Amblyopinus and is closer to that genus than to 
the others of the tribe. Nevertheless, it is more generalized in 
some respects than Amblyopinus; the gula and submentum are 
more like those of Edrabius and Myotyphlus; the labrum is large 
and bilobed; the head capsule is more quediine in form; the elytra 
are longer than broad; the middle tarsi are not spongy beneath. 

Large species; 15 mm. or more in length. Head capsule relatively generalized 
(figs. 30, g; 31, j); almost quediine in form. Eyes large; located near posterior 
margin of head. Head with a very broad neck (fig. 30, g). Labrum large, bilobed 
(fig. 30, g). Gula relatively long and moderately broad (fig. 32, b); gular sutures 
subparallel in part. Elytra longer than broad. Pronotum very large. Hind 
coxae (fig. 33, c) nearly as in Amblyopinus. Legs very strongly spinose. Anterior 
tarsi dilated and spongy beneath; middle tarsi unmodified. 

Megamblyopinus mniszechi Solsky comb. nov. Figures 29, a; 
30, g; 31, ;; 32, b; 33, c; 42, a. 



252 FIELDIANA: ZOOLOGY, VOLUME 37 

Amblyopinus mniszechi Solsky, 1875, Hor. Soc. Ent. Ross., 11: 13, pi. 1, fig. 
4 Hacienda Amable Maria, near La Merced, Junin, Peru. Host un- 
known, suspected of living on "cochins d'Inde (cavias)." 

Length, 17 mm.; maximum width 4.4 mm. Coloration dark reddish-brown. 

Material examined. A female syntype from Fauvel collection, 
Brussels Museum, labeled "Junin, Perou, September, tous un 
coffre." Three specimens, Lago Suche, Moquegua, Peru; 14,600 
feet elevation; collected January 12, 1952, by Oliver P. Pearson; in 
the burrow of Ctenomys opimus nigriceps Thomas. 

Remarks. M. mniszechi and the following species, M. germaini 
Fauvel, are very closely related. The pronotal character used by 
Fauvel to separate the species is of doubtful value, but there are 
diagnostic features. I have examined the specimens that Fauvel 
studied: the type of germaini and a syntype of Solsky's mniszechi. 
In addition, I have studied series of each species that I have com- 
pared with the types. 

M . mniszechi differs from germaini in the structure of the aedeagus 
(fig. 42, a), the sculpture of the head and pronotum, and the darker 
coloration. The lines on the pronotum of M. mniszechi are finer 
and closer than in M. germaini; there are about five lines between 
the punctures of the former species and only about three lines as a 
rule between the punctures of M. germaini. The pronota of the two 
species have the same form. 

The host relationships of Megamblyopinus have some puzzling 
features that need clarification. Jelski first collected M. mniszechi 
under debris in a dwelling and its courtyard and surmised that the 
beetles were associated with domestic guinea pigs. In southern 
Peru Dr. Pearson collected both species of Megamblyopinus in nests 
of tucu-tucus (Ctenomys) and there can be no doubt of their intimate 
association with those rodents. Inasmuch as the Ctenomys of the 
southern provinces of Tacna and Moquegua, Peru, are at the north- 
ern limit of the range of this genus, there can be no doubt that M. 
mniszechi has different host relationships in central Peru. 

Concerning the circumstances under which M. mniszechi was 
collected, Dr. Pearson notes: "Ctenomys opimus was abundant in the 
almost barren, sandy-gravelly desert at Lago Suche. We wounded 
one at a burrow and tried to dig it out; then followed a tunnel for 
about twenty feet to a big stone slab under which a nest was located. 
No guinea pigs and few other mammals occupied abandoned 
Ctenomys burrows in this locality. Because mammal associates 
were so scarce and because the two nests (one at Pichupichuni 






Fig. 42. Aedeagi and apices of aedeagi. a. Megamblyopinus mniszechi Solsky 
b, c. M. germaini Fauvel. 



253 



254 FIELDIANA: ZOOLOGY, VOLUME 37 

containing M. germaini) were practically the only ones we investi- 
gated and yet both had these parasites, I am convinced that the 
beetles were really associated with Clenomys." 

Megamblyopinus germaini Fauvel comb. nov. Figure 42, b, c. 

Amblyopinus germaini Fauvel, 1900, Revue d'Ent., 19: 63 Peru (Fauvel 
collection, Brussels Museum; no host given). 

Length, 16 mm. Color rufous throughout. Distinguished by the characters 
discussed under M. mniszechi. 

Material examined. The type, a female, labeled "Perou." Three 
specimens, Pichupichuni, Puno, Peru; 8 km. northwest of Hua- 
cullani; 12,600 feet elevation; collected April, 1952, by Oliver P. 
Pearson, from a nest of Ctenomys peruanus Sanborn and Pearson. 

Remarks. Dr. Pearson has provided the following notes relating 
to the capture of this species: "Ctenomys peruanus was living in a 
closely cropped grass pampa, heavily grazed by llamas and alpacas. 
We dug out a single Ctenomys nest and found the Megamblyopinus 
specimens there. We found two chambers, one mostly full of 
droppings, the other containing a nest about six inches below the 
surface of the ground. Several mammals made use of old Ctenomys 
burrows, using them after they were abandoned. By far the most 
important of these mammals was a guinea pig, Galea musteloides; 
the others were Galenomys garleppi (an extremely rare mouse at 
the northern limit of its range), Akodon jelskii, and Phyllotis (Aulis- 
comys) boliviensis." 

Genus EDRABIUS Fauvel 

Edrabius Fauvel, 1900, Revue d'Ent., 19: 63. 
Genotype. Edrabius philippianus Fauvel. 

Edrabius differs in a number of important respects from the 
three foregoing genera and was apparently derived from stock that 
diverged from the Amblyopinus line early in tribal evolution. Ed- 
rabius has retained more of the features of the ancestral quediine 
stock; in this respect it resembles the Tasmanian Myotyphlus. 

Head capsule distinctive (figs. 30, b; 31, b); broadly expanded at base so 
that neck is relatively narrow in proportion to head width. Posterior surface of 
head, lateral to neck, concave. Eyes similar to those of Myotyphlus; very small, 
composed of one facet, and located just behind antennal fossae (figs. 30, b; 31, 
b). Labrum large, bilobed (fig. 30, 6). Gula and submentum very similar to those 
of Myotyphlus; gular sutures subparallel, submentum relatively short. Elytra 
subequal in length and width. Hind coxae (fig. 33, b) similar to those of Myo- 



SEEVERS: REVISION OF AMBLYOPININI 255 

typhlus; quediine in form. Middle tarsi of male feebly expanded and spongy; 
without ctenidia. Eighth male sternite emarginate. Lateral plates of ninth 
abdominal segment (fig. 44, g) with a distinctive cluster of extremely long, acicu- 
late setae. 

Edrabius philippianus Fauvel. Figures 30, 6; 31, b; 32, c; 33, b; 

44, a. 

Edrabius philippianus Fauvel, 1900, Revue d'Ent., 19: 65 Antofagasta, 
Chile, in desert of Atacama (Brussels Museum; host, Ctenomys sp.); 
Costa Lima, 1936, Mem. Inst. Oswaldo Cruz, 31: 65. 

It is necessary at present to rely on the female type for our 
concept of E. philippianus inasmuch as none of the Peruvian or 
Argentine specimens that I have examined are conspecific with it. 
It is unfortunate that the aedeagus of this species could not be 
compared with those of the new species herein described. 

The following notes are based on the type female: 

Length, 6 mm.; width of abdomen, 2.5 mm. Coloration reddish-brown. Head 
and pronotum with fine, medium-dense punctation and with fine, very small- 
meshed reticulation. Pronotum with only extremely sparse pubescence on humeri 
and a few hairs close to lateral margins as far back as marginal seta. Elytra with 
sparse clothing of fine, short hairs. Tergites similarly clothed, the hairs longer 
near paratergites. Lateral margin of pronotum as in figure 44, a; the sides con- 
verging strongly from about the middle so that the apex of the pronotum is only 
seven-tenths as broad as the base. 

Material examined. The type, a female, from Antofagasta, 
Chile. 

Edrabius weiseri sp. nov. Figures 43, d; 44, b. 

The largest known species of the genus. E. weiseri differs from 
E. philippianus in the relatively coarse punctation and the obsoles- 
cent reticulation of head and pronotum, the vestiture of elytra and 
abdomen, and the form of the pronotum. 

Length, 9 mm.; width of abdomen, 2.3 mm. Head and pronotum with 
medium-coarse, comparatively dense punctation. Reticulation of head and pro- 
notum obsolescent; in those areas where it is evident the lines form a more coarse- 
meshed reticulation than in E. philippianus. Pronotum with a cluster of moder- 
ately dense, golden pubescence near humeri and along lateral margins nearly to 
base. Elytra with moderately dense vestiture of golden yellow, semi-recumbent 
hairs. Tergites with a similar vestiture except that the hairs of a medium strip 
are finer and less conspicuous. Lateral margin of pronotum as in figure 44, b; 
evenly arcuate so that apex of pronotum is about nine-tenths as broad as base. 
Aedeagus as in figure 43, d. 

Holotype. A male from Troquero, Jujuy, Argentina; collected 
April 1, 1920, by Weiser. Host: no record, almost certainly a 







Fig. 43. Apices of aedeagi. a. Edrabius alticolus sp. nov. b. E. peruanus 
sp. nov. c. E. argentinus sp. nov. d. E. weiseri sp. nov. e. E. pearsoni sp. nov. 



256 












11 w 



".JIlMlll,!!, 

^ .ti/ii.iii 



Fig. 44. Lateral margins of pronota, lateral plates of ninth abdominal seg- 
ment, and eighth male sternites. a. Edrabius philippianus Fauvel. b. E. weiseri 
sp. nov. c, j. E. argentintis sp. nov. d, i. E. pearsoni sp. nov. e. E. alticolus 
sp. nov. f-h. E. peruanus sp. nov. 



257 



258 FIELDIANA: ZOOLOGY, VOLUME 37 

species of Ctenomys. In collection of Chicago Natural History Mu- 
seum. 

Remarks. The type of this species was in the Max Bernhauer 
collection with the manuscript name, E. weiseri n. sp., when the 
collection reached Chicago Natural History Museum. The specimen 
had been sent to Bernhauer by Carlos Bruch and it is very likely 
that other specimens bearing this manuscript name are in the Bruch 
collection in Argentina. 

Edrabius argentinus sp. nov. Figures 43, c; 44, c, j. 

Distinguished from E. philippianus by pronotal form and sculp- 
ture of head and pronotum; from the other new species by its 
distinctive aedeagus. 

Length, 6-7.7 mm.; width of abdomen, 1.7-2.1 mm. Coloration reddish- 
brown. Head and pronotum with fine, relatively sparse punctation; reticulation 
weak to obsolescent, and the fine wavy lines with relatively few cross-connections. 
Pronotum with moderately dense golden pubescence near humeri and along sides. 
Elytra with inconspicuous vestiture of fine, short, recumbent hairs. Tergites 
similarly clothed except that hairs become longer near side margins; the last few 
tergites entirely clothed with yellow hairs. 

Holotype. A male from Hualfin, Catamarca, Argentina (near 
Tucuman); collected December, 1921, by Weiser; "ex nido de 
Ctenomys." In collection of Chicago Natural History Museum. 
The type series is from the Bernhauer collection (ex Carlos Bruch). 

Paratypes. Two specimens, same data as type. 

Edrabius pearsoni sp. nov. Figures 29, b; 43, e; 44, d, i. 

Distinguished from E. philippianus by the form of the pronotum, 
from E. weiseri and E. argentinus by the reticulation of the head 
and pronotum and the form of the aedeagus, and from the other 
Peruvian species by the form of the aedeagus and the eighth male 
sternite. 

Length, 6-6.5 mm.; width of abdomen, 1.7-2.2 mm. Head and pronotum with 
fine, medium-dense punctation and with fine, very small-meshed, sub-hexagonal 
reticulation. Pronotal pubescence very short and sparse near humeri and along 
lateral margins. Elytra and tergites with relatively uniform clothing of fine, 
moderately long hairs. Lateral margin of pronotum as in figure 44, d. Eighth 
male sternite as in figure 44, i; aedeagus as in figure 43, e. 

Holotype. A male from Rio Santa Rosa, Puno, Peru; 13,300 
feet elevation; collected December 15, 1951, by Oliver P. Pearson. 
Host: Ctenomys opimus nigriceps Thomas. In collection of Chicago 
Natural History Museum. 



SEEVERS: REVISION OF AMBLYOPININI 259 

Paratypes. Eighteen specimens, same data as type. 

Edrabius peruanus sp. nov. Figures 43, b; 44, f-h. 

Most closely allied to E. pearsoni; distinguished by form of 
aedeagus, surface contours of pronotum, and minor differences in 
vestiture of elytra and tergites. 

Length, 6.3-7 mm. Color reddish-brown. Head and pronotum with fine, 
medium-dense punctation and with fine, very small-meshed reticulation that 
becomes obsolescent in some areas of pronotal disk. Pronotal pubescence very 
short and sparse in humeral region. Elytra and tergites sparsely and incon- 
spicuously clothed with fine, recumbent hairs. Pronotum (fig. 44, /) with form of 
that of E. pearsoni but with surface depressed medial to humeri. Eighth male 
sternite as in figure 44, h. Aedeagus as in figure 43, b. 

Holotype. A male from Pichupichuni, 8 km. north of Huacullani, 
Puno, Peru; 12,600 feet elevation; collected April, 1952, by Oliver 
P. Pearson. Host: From nest of Ctenomys peruanus Sanborn and 
Pearson. In collection of Chicago Natural History Museum. 

Paratypes. Three specimens, same data as type. Two speci- 
mens, Pampa Queullecota, 60 km. south of Llave, Puno, Peru; 
13,200 feet elevation; collected February 5, 1952, by Oliver P. 
Pearson; from Ctenomys peruanus. 

Edrabius alticolus sp. nov. Figures 43, a; 44, e. 

Distinguished from the two Peruvian species by the form of 
the pronotum and by the aedeagus. 

Length, 5.6-7 mm. Color light reddish-brown. Head and pronotum with 
fine, medium-dense punctation and with fine, very small-meshed, sub-hexagonal 
reticulation. Pronotum with a small patch of yellow pubescence near humeri 
and a few hairs along margin. Elytra and tergites with a uniform vestiture of 
fine yellow hairs. Pronotal form as in figure 44, e. Aedeagus as in figure 43, a. 

Holotype. A male from Pampa de Titire, 29 km. northeast of 
Tarata, Tacna, Peru; 14,600 feet elevation; collected April 13, 1952, 
by Oliver P. Pearson. Host: Ctenomys fulvus Philippi. In collection 
of Chicago Natural History Museum. 

Paratypes. Seven specimens, same data as type. 

Genus MYOTYPHLUS Fauvel 

Myotyphlus Fauvel, 1883, Revue d'Ent., 2: 37; 1900, Revue d'Ent., 19: 61. 
Cryptommatus Matthews, 1884, Cist. Ent., 3: 88. 

Genotype. Myotyphlus jansoni Matthews. 



260 FIELDIANA: ZOOLOGY, VOLUME 37 

Myotyphlus jansoni Matthews. Figures 29, d; 30, a; 31, a; 32, a; 
33, a, /, k, I. 

Amblyopinus jansoni Matthews, 1878, Cist. Ent., 2: 275, pi. 6, figs. 1-8 
Gould's Country, Tasmania (Brit. Mus.; host: "from the fur of a living 
rat"). 

Myotyphlus jansoni Matthews; Fauvel, 1883, Revue d'Ent., 2: 37. 

Cryptommatus jansoni Matthews, 1884, Cist. Ent., 3: 88. 

Myotyphlus jansoni of Tasmania is without doubt the most 
generalized species of Amblyopinini and serves to link the Neo- 
tropical genera with the Quediini (for a discussion of this matter 
see p. 218). 

Head capsule generalized (figs. 30, a; 31, a); quediine in form. Head with a 
medium broad neck inserted in pronotum (fig. 30, a). Gula very long and rela- 
tively narrow (fig. 32, a), the gular sutures parallel; submentum relatively short. 
Labrum large (fig. 30, a). Eyes very small (fig. 31, a) and composed of one facet; 
eyes located just behind antennal fossae. Pronotum quediine in form. Elytra 
longer than broad (fig. 33,/). Hind coxae typically quediine in form (fig. 33, a). 
Anterior tarsi dilated and densely pilose beneath; middle tarsi not dilated and 
pilose but their basal segment with a row of black teeth (ctenidium) along the 
inner margin of the plantar surface. Eighth male sternite feebly emarginate 
(fig. 33, k). Aedeagus with relatively simple lateral lobes (fig. 33, I). 



LIST OF MAMMALIAN HOSTS 

A. Classification of Host Genera 
RODENTIA 

Suborder Myomorpha, Family Cricetidae, Subfamily Cricetinae 
Genera: Oryzomys Baird, Nectomys Peters, Thomasomys Coues, 
Chilomys Thomas, Peromyscus Gloger, Akodon Meyen, 
Oxymycterus Waterhouse, Phyllotis Waterhouse, 
Chinchillula Thomas 

Suborder Hystricomorpha, Family Dasyproctidae, Subfamily Cuniculinae 
Genus Cuniculus Brisson 

Suborder Hystricomorpha, Family Ctenomyidae 
Genus Ctenomys De Blainville 

Suborder Sciuromorpha, Family Heteromyidae 
Genus Heteromys Desmaret 

MARSUPALIA 

Family Didelphidae 

Genera: Didelphis Linnaeus, Metachirus Burmeister, Monodelphis 
Burnett, Marmosa Gray 



B. Host Records 



AMBLYOPINUS Solsky 
jelskii group 
jelskii Solsky 
brandesi Kraatz 
punae sp. nov. 
monticolus sp. nov. 
akodoni sp. nov. 
kofordi sp. nov. 
schmidti Seevers 
colombiae sp. nov. 
piurae sp. nov. 
ancashi sp. nov. 
sp. 

spinipennis sp. nov. 
sp. 
longus Franz 

fuegensis group 
fuegensis Arrow 

hershkovitzi group 
hershkovitzi sp. nov. 



Peru 

Peru 

Peru 

Peru 

Peru 

Peru 

Guatemala 

Colombia 

Peru 

Peru 

Peru 

Peru 

Peru 

Brazil 



Chile 



Colombia 



(l)Oryzomys laticeps Lund 
No record 

Akodon jelskii pulcherrimus Thomas 
Ctenomys opimus nigriceps Thomas 
Akodon jelskii pyrrhotis Thomas 
Akodon berlepschii Thomas 
Peromyscus guatemalensis Merriam 
Thomasomys aureus nicefori Thomas 
Thomasomys sp. 
Phyllotis darwini Waterhouse 
Phyllotis sublimis Thomas 
Akodon aerosus Thomas 
Chinchillula sahamae Thomas 
Oxymycterus rufus Desmaret 
Akodon (Thaptomys) nigrita Licht. 

Ctenomys magellanicus dicki Osgood 



Heteromys anomalus Thompson 



261 



262 



FIELDIANA: ZOOLOGY, VOLUME 37 



sanborni group 




sanborni Seevers 


Peru 


emarginatus sp. nov. 


Colombia 


chilomysi sp. nov. 


Colombia 


montivagans sp. nov. 


Colombia 


oryzomysi sp. nov. 


Colombia 


huilae sp. nov. 


Colombia 


waterhousei group 




waterhousei Fauvel 


Ecuador, 




Colombia 


kalinowskii sp. nov. 


Peru 


pacae sp. nov. 


Peru 


sp. 


Peru 


henseli group 




henseli Kolbe 


Brazil 



bequaerti Notman 
marmosae Seevers 

Incertae sedis 

angustus Arrow 



Brazil 
Brazil 



Thomasomys sp. 
(l)Carollia perspicillata L. 
Oryzomys keaysi Allen 
Thomasomys laniger Thomas 
Chilomys sp. 
Chilomys sp. 

Oryzomys albigularis Tomes 
Chilomys sp. 



Didelphis albiventris Lund 
Didelphis albiventris Lund 
Cuniculus taczanowskii Stolzmann 
Cuniculus taczanowskii Stolzmann 



Metachirus opossum Linnaeus 
Didelphis marsupialis Linnaeus 
Monodelphis sp. 
Marmosa incana paulensis Tate 



British Guiana No record 



AMBLYOPINODES gen. nov. 

gahani Fauvel Argentina 

piceus Brethes 

claviger Franz 
travassosi Costa Lima 

peruvianas sp. nov. 
similis sp. nov. 



Argentina, 
Brazil 
Brazil 
Brazil 

Peru 
Peru 



No record 

Nectomys squamipes Brants 
Oxymycterus rufus Desmaret 
Thaptomys nigrita Lichtenstein 
Oxymycterus judex Thomas 
Oryzomys sp. 
Oxymycterus juliacae Allen 



MEGAMBLYOPINUS gen. nov. 

mniszechi Solsky Peru (Junin) 

Peru 
(Moquegua) 

germaini Fauvel Peru 



No record 

Ctenomys opimus nigriceps Thomas 
Ctenomys peruanus Sanborn and 
Pearson 



EDRABIUS Fauvel 



philippianus Fauvel 


Chile 


weiseri sp. nov. 


Argentina 


argentinus sp. nov. 


Argentina 


pearsoni sp. nov. 


Peru 


peruanus sp. nov. 


Peru 


alticolus sp. nov. 


Peru 



Ctenomys sp. 

(l)Ctenomys sp. 

Ctenomys sp. 

Ctenomys opimus nigriceps Thomas 

Ctenomys peruanus Sanborn and 

Pearson 

Ctenomys fulvus Philippi 



MYOTYPHLUS Fauvel 
jansoni Matthews Tasmania 



A "rat" 



REFERENCES 

Arrow, G. J. 

1907. On two new parasitic Coleoptera (family Staphylinidae) from South 
America. Ann. Mag. Nat. Hist., (7), 19: 125-127. 

Brethes, J. 

1926. Un nouveau staphylin (Col.) muricole de la Republique Argentina. 
An. Mus. Nac. Hist. Nat. Bernardino Rivadavia, 34: 17-20. 

Costa Lima, Angelo M. da 

1927a. Contribucao ao estudo de coleopteros staphylinideos encontrados no 

pello de murideos. C. R. Soc. Biol., 79: 842. 
1927b. Contribucao ao estudo de coleopteros staphylinideos encontrados no 

pello de murideos. Sci. Med. Rio Janeiro, 5: 380-383. 
1936. Sobre os generos Amblyopinus e Edrabius. Mem. Inst. Oswaldo Cruz, 

31: 55-68, 1 pi., 4 text figs. 

Fauvel, Albert 

1883. Amblyopinus et My otyphlus. Revue d'Ent., 2: 37-40. 

1900. Amblyopinus, Myotyphlus et Edrabius. Revue d'Ent., 19: 61-66. 

1901. Amblyopinus nouveau. Revue d'Ent., 20: 5-6. 

FONSECA, FLAVIO DA 

1939. Especies de Amblyopinus parasitas de murideos e didelfideos em S. 
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Franz, Elli 

1927. Bemerkungen zur Gattung Amblyopinus Solsky. Abh. Senckenb. 
Naturf. Ges. Frankfurt a/M., 40: 405-408, 4 figs. 

1930. Zwei neue Vertreter der Gattung Amblyopinus Solsky (Ins. Col.). 
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Kolbe, H. J. 

1911. Uber ekto- und endoparasitische Coleopteren. Deut. Ent. Nat. Bibl., 
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Kraatz, G. 

1900. Amblyopinus brandesi n. sp. Deut. Ent. Zeitschr., 1900: 212. 

Matthews, A. 

1878. On the genus Amblyopinus, and description of a new species from 
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1884. Notes on M. Fauvel 's observations on Amblyopinus jansoni, with a 
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10 figs. 

NOTMAN, H. 

1923. A new genus and species of Staphylinidae parasitic on a South American 
opossum. Amer. Mus. Nov., no. 68, 3 pp. 

263 



264 FIELDIANA: ZOOLOGY, VOLUME 37 

Seevers, Charles H. 

1944. A new subfamily of beetles parasitic on mammals. Staphylinidae, Am- 
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Solsky, Simon M. 

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1951. Evolution of the early Tertiary rodents. Evolution, 5: 207-215. 

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1939. Amblyopinus henseli Kolbe, um coleoptero da familia Staphylinidae que 
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